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For Peer Review Biological Journal of the Linnean Society Marine tethysuchian c rocodyliform from the ?Aptian -Albian (Early Cretaceous) of the Isle of Wight, England Journal:For Biological Peer Journal of theReview Linnean Society Manuscript ID: BJLS-3227.R1 Manuscript Type: Research Article Date Submitted by the Author: 05-May-2014 Complete List of Authors: Young, Mark; University of Edinburgh, Biological Sciences; University of Southampton, School of Ocean and Earth Science Steel, Lorna; Natural History Museum, Earth Sciences Foffa, Davide; University of Bristol, Department of Earth Sciences Price, Trevor; Dinosaur Isle Museum, Naish, Darren; University of Southampton, School of Ocean and Earth Science Tennant, Jon; Imperial College London, Department of Earth Science and Engineering Albian, Aptian, Cretaceous, Dyrosauridae, England, Ferruginous Sands Keywords: Formation, Isle of Wight, Pholidosauridae, Tethysuchia, Upper Greensand Formation Biological Journal of the Linnean Society Page 1 of 50 Biological Journal of the Linnean Society 1 2 3 Marine tethysuchian crocodyliform from the ?Aptian-Albian (Early Cretaceous) 4 5 6 of the Isle of Wight, England 7 8 9 10 by MARK T. YOUNG 1,2 *, LORNA STEEL 3, DAVIDE FOFFA 4, TREVOR PRICE 5 11 12 2 6 13 DARREN NAISH and JONATHAN P. TENNANT 14 15 16 1 17 Institute of Evolutionary Biology, School of Biological Sciences, The King’s Buildings, University 18 For Peer Review 19 of Edinburgh, Edinburgh, EH9 3JW, United Kingdom 20 21 2 School of Ocean and Earth Science, National Oceanography Centre, University of Southampton, 22 23 Southampton, SO14 3ZH, United Kingdom 24 25 3 Department of Earth Sciences, Natural History Museum, London, SW7 5BD, United Kingdom 26 27 4 School of Earth Sciences, Wills Memorial Building, University of Bristol, Bristol, BS8 1RJ, United 28 29 Kingdom 30 31 5 Dinosaur Isle Museum, Sandown, Isle of Wight, PO36 8QA, United Kingdom 32 33 6 Department of Earth Science and Engineering, Imperial College London, London, SW6 34 35 2AZ, United Kingdom 36 37 38 39 40 * Corresponding author: [email protected] 41 42 43 44 RUNNING HEAD: Cretaceous tethysuchian from England 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Biological Journal of the Linnean Society Biological Journal of the Linnean Society Page 2 of 50 1 2 3 ABSTRACT. A marine tethysuchian crocodyliform from the Isle of Wight, most likely from 4 5 the Upper Greensand Formation (late Albian, Early Cretaceous), is described. However, we 6 7 cannot preclude it being from the Ferruginous Sands Formation (late Aptian), or more 8 9 10 remotely, the Sandrock Formation (late Aptian-early Albian). The specimen consists of the 11 12 anterior region of the right dentary, from the tip of the dentary to the incomplete fourth 13 14 alveolus. This specimen increases the known geological range of marine tethysuchians back 15 16 into the late Early Cretaceous. While we refer it to Tethysuchia incertae sedis , there are seven 17 18 For Peer Review 19 anterior dentary characteristics which suggest a possible relationship with the Maastrichtian- 20 21 Eocene clade Dyrosauridae. We also review 'middle' Cretaceous marine tethysuchians, 22 23 including putative Cenomanian dyrosaurids. We conclude that there is insufficient evidence 24 25 to be certain that any known Cenomanian specimen can be safely referred to Dyrosauridae, as 26 27 there are some cranial similarities between basal dyrosaurids and Cenomanian-Turonian 28 29 30 marine 'pholidosaurids'. Future study of middle Cretaceous tethysuchians could help unlock 31 32 the origins of Dyrosauridae and improve our understanding of tethysuchian 33 34 macroevolutionary trends. 35 36 37 38 39 Key words: Albian – Aptian – Cretaceous – England – Tethysuchia – Upper Greensand 40 41 Formation. 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Biological Journal of the Linnean Society Page 3 of 50 Biological Journal of the Linnean Society 1 2 3 INTRODUCTION 4 5 6 Tethysuchian crocodyliforms were a highly successful group, some of which returned 7 8 to a marine lifestyle during the latter part of the Mesozoic and early Cenozoic. Many species 9 10 superficially resembled extant gharials in having enlarged supratemporal fenestrae, an 11 12 elongate, tubular snout and a high tooth count (e.g. Koken, 1887; Mook 1933, 1934; Wu et 13 14 al ., 2001; Jouve et al . 2005a, 2006a; Barbosa et al ., 2008). Tethysuchia was one of several 15 16 17 crocodyliform clades that survived the end-Cretaceous mass extinction event, with the 18 For Peer Review 19 subclade Dyrosauridae continuing to radiate and diversify during the Paleocene and Eocene 20 21 (e.g. Buffetaut 1976, 1978, 1982; Jouve 2005, 2007; Jouve et al . 2005a, 2005b, 2006a, 2008; 22 23 Barbosa et al ., 2008; Hill et al ., 2008; Hastings et al . 2010, 2011). However, the origins of 24 25 26 this clade are poorly understood due to a paucity of fossils from the ‘middle’ Cretaceous. 27 28 A phylogenetic definition of Tethysuchia was recently proposed by Andrade et al . 29 30 (2011: S102) as: "the clade composed of Pholidosaurus purbeckensis (Mansell-Pleydell, 31 32 1888) and Dyrosaurus phosphaticus (Thomas, 1893), their common ancestor and all its 33 34 descendants". This definition encompasses Dyrosauridae and Pholidosauridae, and possibly 35 36 37 Elosuchidae. Although curiously, this is not the case in the phylogenetic analysis of Andrade 38 39 et al . (2011), as Elosuchidae was recovered outside the Pholidosaurus + Dyrosaurus clade. 40 41 Moreover, Andrade et al . (2011) proposed that Elosuchidae be used for the clade consisting 42 43 of Elosuchus , Sarcosuchus and Vectisuchus . Also, a phylogenetic definition of 44 45 46 Pholidosauridae was recently proposed by Fortier et al . (2011: S259) as: "a stem-based group 47 48 name including Pholidosaurus schaumburgensis (Meyer, 1841) and all taxa closer to it than 49 50 to Dyrosaurus phosphaticus (Thomas, 1893) or Pelagosaurus typus Bronn, 1841". No 51 52 explicit phylogenetic definition has been proposed for Dyrosauridae or Elosuchidae. 53 54 Phylogenetic analyses consistently finds Dyrosauridae to be holophyletic (e.g. Wu et 55 56 57 al. , 2001; Jouve, 2005; Jouve et al . 2005a, 2006a, 2008; Barbosa et al ., 2008; Young & 58 59 60 Biological Journal of the Linnean Society Biological Journal of the Linnean Society Page 4 of 50 1 2 3 Andrade, 2009; Hastings et al . 2010, 2011; Andrade et al ., 2011; Fortier et al ., 2011). The 4 5 holophyly of Pholidosauridae however, is not always recovered. Pholidosauridae has either 6 7 been found to be a paraphyletic grade of taxa closely related to Dyrosauridae (Wu et al. , 8 9 10 2001; Jouve et al . 2005a, 2008; Barbosa et al ., 2008; Young & Andrade, 2009; Hastings et al . 11 12 2010, 2011; Andrade et al ., 2011), holophyletic (Fortier et al ., 2011), or holophyletic with 13 14 Elosuchus being outside the clade comprising Dyrosauridae and Pholidosauridae (Jouve et 15 16 al ., 2006a). Martin & Buffetaut (2012) and Martin et al . (in press) also found Pholidosauridae 17 18 For Peer Review 19 to be holophyletic, but as no dyrosaurids were included in those analyses they never tested 20 21 whether pholidosaurids constitute a natural group. When Pholidosauridae is found to be 22 23 paraphyletic, the sister taxon of Dyrosauridae varies between Terminonaris (Wu et al. , 2001; 24 25 Jouve et al . 2005a, 2008; Barbosa et al ., 2008), Oceanosuchus (Young & Andrade, 2009) and 26 27 Elosuchus (Hastings et al . 2010, 2011); although Oceanosuchus was only included in the 28 29 30 analysis of Young & Andrade (2009). As such, the internal evolutionary relationships of 31 32 Tethysuchia are in flux. This has important implications for tethysuchian evolution, in 33 34 particular for our understanding of dyrosaurid origins, and whether dyrosaurids and the 35 36 marine 'pholidosaurids' Terminonaris and Oceanosuchus constitute a single marine radiation 37 38 39 or several independent ones. 40 41 One of the major issues hampering resolution in these analyses is the paucity of 42 43 'middle' Cretaceous (Barremian-Turonian) marine tethysuchians, thus affecting our 44 45 understanding of character polarity. In those phylogenetic analyses where 'pholidosaurids' are 46 47 paraphyletic, all potential dyrosaurid sister taxa ( Elosuchus , Oceanosuchus and 48 49 50 Terminonaris ) are from this time span (see Mook 1933, 1934; de Lapparent de Broin, 2002; 51 52 Wu et al ., 2001; Hua et al ., 2007). Furthermore, the earliest potential dyrosaurids are 53 54 Cenomanian in age (Buffetaut & Lauverjat, 1978; Buffetaut et al ., 1990). Therefore, 55 56 57 58 59 60 Biological Journal of the Linnean Society Page 5 of 50 Biological Journal of the Linnean Society 1 2 3 investigating specimens from the Barremian-Turonian stages will be key to elucidating the 4 5 early evolution of Tethysuchia. 6 7 Here we describe a long known but previously unstudied tethysuchian crocodyliform. 8 9 10 This specimen, the anterior-most part of a right dentary (NHMUK PV OR36173), is most 11 12 likely from the Upper Greensand Formation of England (late Albian, Early Cretaceous). 13 14 Although this specimen was discovered over 150 years ago, it has only been briefly 15 16 mentioned once in the literature. Furthermore, it is of importance due to its unusual 17 18 For Peer Review 19 morphology and geological age. The presence of a marine tethysuchian in the late Early 20 21 Cretaceous of England would indicate that tethysuchians moved into the marine realm earlier 22 23 than previously realised. 24 25 26 27 28 ABBREVIATIONS 29 30 INSTITUTIONAL 31 32 MIWG, the Museum of Isle of Wight Geology (now IWCMS- the Isle of Wight County 33 34 Museum Service, incorporating Dinosaur Isle museum and visitor attraction); NHMUK, 35 36 37 Natural History Museum, London, England, United Kingdom. 38 39 40 41 ANATOMICAL 42 43 D1, first dentary alveolus; D2, second dentary alveolus; D3, third dentary alveolus; D4, 44 45 46 fourth dentary alveolus; for, foramen; rug, rugose patch; sym, symphysis.
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